EP0257542B1 - Resistenzgen gegen Phosphinothricin und seine Verwendung - Google Patents

Resistenzgen gegen Phosphinothricin und seine Verwendung Download PDF

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Publication number
EP0257542B1
EP0257542B1 EP87112023A EP87112023A EP0257542B1 EP 0257542 B1 EP0257542 B1 EP 0257542B1 EP 87112023 A EP87112023 A EP 87112023A EP 87112023 A EP87112023 A EP 87112023A EP 0257542 B1 EP0257542 B1 EP 0257542B1
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EP
European Patent Office
Prior art keywords
ptc
resistance
ptt
gene
fragment
Prior art date
Legal status (The legal status is an assumption and is not a legal conclusion. Google has not performed a legal analysis and makes no representation as to the accuracy of the status listed.)
Expired - Lifetime
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EP87112023A
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German (de)
English (en)
French (fr)
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EP0257542A2 (de
EP0257542A3 (en
Inventor
Eckhard Strauch
Wolfgang Dr. Wohlleben
Walter Arnold
Renate Alijah
Alfred Prof. Dr. Pühler
Gerhard Dr. Wöhner
Rüdiger Dr. Marquardt
Susanne Dr. Grabley
Dieter Dr. Brauer
Klaus Dr. Bartsch
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Bayer CropScience AG
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Hoechst AG
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Filing date
Publication date
Priority claimed from DE19863628747 external-priority patent/DE3628747A1/de
Priority claimed from DE19863642829 external-priority patent/DE3642829A1/de
Priority claimed from DE19873700313 external-priority patent/DE3700313A1/de
Application filed by Hoechst AG filed Critical Hoechst AG
Publication of EP0257542A2 publication Critical patent/EP0257542A2/de
Publication of EP0257542A3 publication Critical patent/EP0257542A3/de
Application granted granted Critical
Publication of EP0257542B1 publication Critical patent/EP0257542B1/de
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    • C—CHEMISTRY; METALLURGY
    • C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
    • C12P—FERMENTATION OR ENZYME-USING PROCESSES TO SYNTHESISE A DESIRED CHEMICAL COMPOUND OR COMPOSITION OR TO SEPARATE OPTICAL ISOMERS FROM A RACEMIC MIXTURE
    • C12P41/00—Processes using enzymes or microorganisms to separate optical isomers from a racemic mixture
    • C12P41/006—Processes using enzymes or microorganisms to separate optical isomers from a racemic mixture by reactions involving C-N bonds, e.g. nitriles, amides, hydantoins, carbamates, lactames, transamination reactions, or keto group formation from racemic mixtures
    • C12P41/007—Processes using enzymes or microorganisms to separate optical isomers from a racemic mixture by reactions involving C-N bonds, e.g. nitriles, amides, hydantoins, carbamates, lactames, transamination reactions, or keto group formation from racemic mixtures by reactions involving acyl derivatives of racemic amines
    • C—CHEMISTRY; METALLURGY
    • C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
    • C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
    • C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
    • C12N15/09—Recombinant DNA-technology
    • C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
    • C12N15/64—General methods for preparing the vector, for introducing it into the cell or for selecting the vector-containing host
    • C—CHEMISTRY; METALLURGY
    • C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
    • C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
    • C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
    • C12N15/09—Recombinant DNA-technology
    • C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
    • C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
    • C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
    • C12N15/8201—Methods for introducing genetic material into plant cells, e.g. DNA, RNA, stable or transient incorporation, tissue culture methods adapted for transformation
    • C12N15/8209—Selection, visualisation of transformants, reporter constructs, e.g. antibiotic resistance markers
    • C—CHEMISTRY; METALLURGY
    • C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
    • C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
    • C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
    • C12N15/09—Recombinant DNA-technology
    • C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
    • C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
    • C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
    • C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
    • C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
    • C12N15/8271—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance
    • C12N15/8274—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield for stress resistance, e.g. heavy metal resistance for herbicide resistance
    • C12N15/8277—Phosphinotricin
    • C—CHEMISTRY; METALLURGY
    • C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
    • C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
    • C12N9/00—Enzymes; Proenzymes; Compositions thereof; Processes for preparing, activating, inhibiting, separating or purifying enzymes
    • C12N9/10—Transferases (2.)
    • C12N9/1025—Acyltransferases (2.3)
    • C12N9/1029—Acyltransferases (2.3) transferring groups other than amino-acyl groups (2.3.1)

Definitions

  • PTC Phosphinothricin
  • PTC 2-amino-4-methylphosphinobutyric acid
  • PTC is a glutamine synthetase inhibitor.
  • PTC is a "building block" of the antibiotic phosphinothricyl-alanyl-alanine.
  • This tripeptide is active against Gram-positive and Gram-negative bacteria and also against the fungus Botrytis cinerea (Bayer et al., Helv. Chim. Acta 55 (1972) 224).
  • PTT is produced by the strain Streptomyces viridochromogenes Tü 494 (DSM 40736, DSM 4112).
  • the invention which is defined in the patent claims, relates to a resistance gene against PTC and its use for producing PTC-resistant plants.
  • This gene can also be used as a resistance marker.
  • the gene is also suitable for the selective N-acetylation of the L-form of racemic PTC.
  • the resistance gene according to the invention against PTC can be obtained from the total DNA of Streptomyces viridochromogenes DSM 4112 selected for PTT resistance by cutting with Bam HI, cloning a 4.0 kb fragment and selection for PTT resistance.
  • the restriction map (FIG. 1) characterizes this 4.0 kb fragment in more detail.
  • the position of the coding area was narrowed further by cloning partial areas of this 4 kb fragment. It was found here that the resistance gene is located on the 1.6 kb Sst II- Sst I fragment (positions 0.55 to 2.15 in FIG. 1). The 0.8 kb fragment is obtained by digestion with Bgl II, which after incorporation into a plasmid and transformation of S. lividans PTT resistance mediated. This resistance is due to the N-acetylation of PTC.
  • DNA sequence I (Appendix).
  • the position of the resistance gene can be determined from the open reading frames of this sequence (from position 258).
  • the end of the gene is at the penultimate of the nucleotides shown (position 806), i. H. the last nucleotide (position 807) is the start of the stop codon.
  • DNA sequence I the "Shine-Dalgarno sequence" is highlighted by underlining, as is the GTG acting as the start codon. The relevant reading frame is therefore printed on the top line.
  • DNA sequence II shows the restriction sites within the sequenced gene. Enzymes that cut the sequence more than six times are not specified.
  • the antibiotic PTT is absorbed by bacteria and broken down to PTC. This also inhibits glutamine synthetase in bacteria, so that the bacteria die from a lack of glutamine. PTT-producing bacteria should therefore have a mechanism that protects them from the effects of the PTT, that is, either prevents the resumption of the PTT produced or enables a modification of the PTC degradation product.
  • the PTT producer S. viridochromogenic DSM 4112 but sensitive to its own antibiotic. Unexpectedly, however, by selecting for PTT resistance at the surprisingly high rate of about 10 ⁇ 5 selectants that are resistant to PTT and also suppress the underground growth of the neighboring colonies.
  • a gene bank was created from the DNA of these selectants by isolating the DNA, cleaving with Bam HI and ligating into a streptomycete vector.
  • the ligation mixture was in the commercially available strain S. lividans TK 23 transformed, each 1 ⁇ g ligation mixture about 5000 to 10000 transformants with an insert of about 1 to 5 kb were obtained. PTT-resistant S are found among the transformants. lividans strains. By isolation of the plasmid and retransformation in S. lividans could be shown that the resistance is plasmid-encoded. The gene responsible for resistance is located on a 4 kb Bam HI fragment (FIG. 1).
  • the coding area is located on the 0.8 kb Bgl II fragment.
  • the Bam HI fragment contains no interfaces for the enzymes Cla I, Eco RI, Eco RV, Hin dIII, Hpa I, Kpn I, Pvu I, Pvu II and Xho I.
  • the resistance gene according to the invention can thus be transformed into plants with suitable vectors after coupling to plant promoters, and PTC-resistant plants can thus be produced.
  • N-acetylation of PTC can also be used for the racemate separation of synthetic D, L-PTC, since only the L-form is selectively acetylated.
  • the invention thus also relates to the use of the resistance gene for the selective N-acetylation of the L form of racemic PTC.
  • the PTCAcetyltransferase encoded by the resistance gene according to the invention can thus be used to separate racemic PTC, such as is obtainable, for example, from German Patent No. 2,717,440, into the optical antipodes by exposing the racemate to the acetylating action of this enzyme, the selectively L-shape is attacked while the D-shape remains unchanged.
  • the mixture obtained in this way can then be separated in a manner known per se on account of its different properties.
  • N-acyl-D, L-amino acids It is known to bring N-acyl-D, L-amino acids into contact with optionally fixed acylases, selectively releasing the L-amino acid from the mixture with the N-acyl-D-amino acid after acidification with water-immiscible solvents can be extracted (British Patent 1,369,462).
  • a corresponding separation of N-acyl-D, L-PTC is known for example from German Offenlegungsschrift 2 939 269 or US Pat. No. 4,226,941.
  • the D-PTC remaining according to the invention can be racemized in a known manner (European patent application with publication number (EP-A) 0 137 371, example 8) and then returned to the process.
  • the isolation of the enzyme which is to be understood here and in the following also the enzymatically active part, is possible, but not necessary. If the enzyme is isolated, it can be used in free or carrier-fixed form. Suitable carriers are described, for example, in EP-A 0 141 223. However, the enzyme is expediently not isolated, instead any PTC-resistant cells are used which express the enzyme according to the invention.
  • the PTT-resistant selectant of S. viridochromogenic DSM 4112 can be used. Any cell which is transformed with the gene according to the invention and which is able to express the PTC acetyltransferase can also advantageously be used.
  • the gene according to the invention which also includes active parts thereof, can be introduced into the host cell in a plasma-integrated form or with other conventional genetic engineering methods, for example by transfection.
  • the cells which express the PTC acetyltransferase can be used in free or fixed form, the usual methods of fixation being used (e.g. German Offenlegungsschrift 3,237,341 and the literature cited therein).
  • the enzymatic acetylation of L-PTC according to the invention takes place in the manner customary for enzymatic reactions, whereby the process conditions are based on the conditions of the organism used. Basically, the same methods can be used as for the selective deacylation processes mentioned above.
  • the strain S. viridochromogenic DSM 4112 was grown on minimal medium (Hopwood et al., Genetic Manipulation of Streptomyces, A Laboratory Manual, The John Innes Foundation, Norwich, England (1985), p. 233) and PTT in increasing concentrations was added. At a concentration of 100 ⁇ g / ml a resistant colony was found for every 105 colonies.
  • the plasmid pSVH1 (European patent specification 0 070 522) is cut with Bgl II, the approximately 7.1 kb fragment is isolated and with the 1.1 kb Bcl I fragment with thiostrepton resistance (European patent application with publication number 0 158 201 ) ligated.
  • the 8.15 kb plasmid pEB2 is obtained (FIG. 2).
  • the total DNA is isolated from the selectants according to Example 1 and cleaved with Bam HI.
  • the plasmid pEB2 is also opened with Bam HI, the two batches are combined and ligated.
  • the ligation mixture is according to S. lividans TK 23 (available from the John Innes Foundation), whereby 1 ⁇ g of ligation mixture contains 5,000 to 10,000 transformants with an insert of about 1-5 kb be preserved. Selection for PTT resistance results in 2 resistant S. lividans colonies.
  • the plasmid taken up is isolated from these and cut with Bam HI. A 4 kb Bam HI fragment is found, which carries the gene responsible for resistance. This plasmid was given the designation pPRl (FIG. 3).
  • strains were examined to demonstrate the acetylating activity of the cloned fragment: S. viridochromogenic DSM 40736, p . viridochromogenes (PTT-resistant mutant), S. lividans TK23 and S. lividans TK 23 (pPR1).
  • the strains are inoculated in lysis medium A (European patent application with publication number 0 158 872, p. 6) and incubated for 2 days at 30 ° C. in a rotary shaker. After harvesting, 1 mg of mycelium is disrupted with ultrasound in a suitable buffer (e.g. RS buffer: CJ Thompson et al., J. Bacteriol. 151 (1982), 678-685).
  • a suitable buffer e.g. RS buffer: CJ Thompson et al., J. Bacteriol. 151 (1982), 678-685.
  • a typical experiment to measure PTC degradation is as follows: 250 ⁇ l of crude extract are mixed with 100 ⁇ l of PTC solution (250 ⁇ g / ml) and 50 ⁇ l of acetyl-CoA (4 mg / ml) and incubated at 30 ° C for 2 hours.

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  • General Chemical & Material Sciences (AREA)
  • Micro-Organisms Or Cultivation Processes Thereof (AREA)
  • Enzymes And Modification Thereof (AREA)
  • Preparation Of Compounds By Using Micro-Organisms (AREA)
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  • Steroid Compounds (AREA)
  • Measuring Or Testing Involving Enzymes Or Micro-Organisms (AREA)
  • Medicines That Contain Protein Lipid Enzymes And Other Medicines (AREA)
  • Transition And Organic Metals Composition Catalysts For Addition Polymerization (AREA)
  • Macromolecular Compounds Obtained By Forming Nitrogen-Containing Linkages In General (AREA)
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EP87112023A 1986-08-23 1987-08-19 Resistenzgen gegen Phosphinothricin und seine Verwendung Expired - Lifetime EP0257542B1 (de)

Applications Claiming Priority (8)

Application Number Priority Date Filing Date Title
DE3628747 1986-08-23
DE19863628747 DE3628747A1 (de) 1986-08-23 1986-08-23 Resistenzgen gegen phosphinothricin und seine verwendung
DE3637307 1986-11-03
DE3637307 1986-11-03
DE3642829 1986-12-16
DE19863642829 DE3642829A1 (de) 1986-08-23 1986-12-16 Resistenzgen gegen phosphinothricin
DE3700313 1987-01-08
DE19873700313 DE3700313A1 (de) 1986-08-23 1987-01-08 Verwendung eines resistenzgens gegen phosphinothricin

Publications (3)

Publication Number Publication Date
EP0257542A2 EP0257542A2 (de) 1988-03-02
EP0257542A3 EP0257542A3 (en) 1990-03-07
EP0257542B1 true EP0257542B1 (de) 1992-05-06

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EP87112023A Expired - Lifetime EP0257542B1 (de) 1986-08-23 1987-08-19 Resistenzgen gegen Phosphinothricin und seine Verwendung

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EP (1) EP0257542B1 (enrdf_load_stackoverflow)
JP (4) JPH0797994B2 (enrdf_load_stackoverflow)
CN (2) CN1040772C (enrdf_load_stackoverflow)
AT (1) ATE75776T1 (enrdf_load_stackoverflow)
AU (1) AU604743B2 (enrdf_load_stackoverflow)
CA (1) CA1337597C (enrdf_load_stackoverflow)
DK (1) DK175254B1 (enrdf_load_stackoverflow)
ES (1) ES2038631T3 (enrdf_load_stackoverflow)
FI (1) FI100251B (enrdf_load_stackoverflow)
GR (1) GR3005200T3 (enrdf_load_stackoverflow)
HU (1) HU217208B (enrdf_load_stackoverflow)
IL (1) IL83604A (enrdf_load_stackoverflow)
NZ (1) NZ221526A (enrdf_load_stackoverflow)

Families Citing this family (31)

* Cited by examiner, † Cited by third party
Publication number Priority date Publication date Assignee Title
DE3765449D1 (de) * 1986-03-11 1990-11-15 Plant Genetic Systems Nv Durch gentechnologie erhaltene und gegen glutaminsynthetase-inhibitoren resistente pflanzenzellen.
CN87100603A (zh) * 1987-01-21 1988-08-10 昂科公司 抗黑素瘤疫苗
DE3716309A1 (de) * 1987-05-15 1988-11-24 Hoechst Ag Resistenzgen gegen phosphinothricin
DE3732972A1 (de) * 1987-07-02 1989-01-12 Hoechst Ag Resistenzgen gegen phosphinothricin und seine verwendung
DE4126414A1 (de) * 1991-08-09 1993-02-11 Hoechst Ag Deacetylasegene zur erzeugung von phosphinothricin oder phosphinothricyl-alanyl-alanin, verfahren zu ihrer isolierung und ihre verwendung
DE4222407C1 (de) * 1992-07-08 1993-10-07 Max Planck Gesellschaft Modulartiges Promotor-Konstrukt
AR008158A1 (es) * 1996-09-05 1999-12-09 Syngenta Participations Ag Proceso para el control de malas hierbas en cultivos de plantas utiles que son resistentes a un fosfo-herbicida y una composicion herbicida para dicho uso.
US6586367B2 (en) 1996-09-05 2003-07-01 Syngenta Crop Protection, Inc. Process for the control of weeds
DE19836659A1 (de) 1998-08-13 2000-02-17 Hoechst Schering Agrevo Gmbh Herbizide Mittel für tolerante oder resistente Baumwollkulturen
DE19836700A1 (de) 1998-08-13 2000-02-17 Hoechst Schering Agrevo Gmbh Herbizide Mittel für tolerante oder resistente Getreidekulturen
DE19836660A1 (de) 1998-08-13 2000-02-17 Hoechst Schering Agrevo Gmbh Herbizide Mittel für tolerante oder resistente Sojakulturen
DE19836684A1 (de) 1998-08-13 2000-02-17 Hoechst Schering Agrevo Gmbh Herbizide Mittel für tolerante oder resistente Reiskulturen
PL212110B1 (pl) * 1998-08-13 2012-08-31 Bayer Cropscience Ag Zastosowanie kompozycji herbicydów w uprawach kukurydzy i sposób zwalczania szkodliwych roślin w uprawach kukurydzy
UA82665C2 (uk) 2002-02-26 2008-05-12 Сингента Лимитед Спосіб вибірного одержання рослин з чоловічою або жіночою стерильністю
WO2006054458A1 (ja) 2004-11-17 2006-05-26 Hokko Chemical Industry Co., Ltd. 除草剤耐性遺伝子及びその利用
KR100743611B1 (ko) * 2006-03-28 2007-08-01 최광술 산업용 로봇의 케이블 조절장치
AR074941A1 (es) 2009-01-07 2011-02-23 Bayer Cropscience Sa Plantas transplastomicas exentas de marcador de seleccion
UY33139A (es) 2009-12-23 2011-07-29 Bayer Cropscience Ag Plantas tolerantes a herbicidas inhibidores de las hppd
CN102762725A (zh) 2009-12-23 2012-10-31 拜尔知识产权有限公司 耐受hppd抑制剂型除草剂的植物
EA201290559A1 (ru) 2009-12-23 2013-01-30 Байер Интеллектуэль Проперти Гмбх Растения, устойчивые к гербицидам - ингибиторам hppd
CN102762724A (zh) 2009-12-23 2012-10-31 拜尔知识产权有限公司 对hppd抑制剂型除草剂耐受的植物
WO2011076892A1 (en) 2009-12-23 2011-06-30 Bayer Cropscience Ag Plants tolerant to hppd inhibitor herbicides
BR112012029616A2 (pt) 2010-05-21 2015-10-20 Bayer Ip Gmbh agentes herbicidas para as culturas de cereais tolerantes ou resistentes
CN103025167A (zh) 2010-05-21 2013-04-03 拜耳知识产权有限责任公司 用于耐受性或抗性稻栽培种的除草剂
BR112012029621A2 (pt) 2010-05-21 2015-09-22 Bayer Ip Gmbh agentes herbicidas para as culturas de milho tolerantes ou resistentes
CN103002743A (zh) 2010-05-21 2013-03-27 拜耳知识产权有限责任公司 用于耐药性或抗药性油菜作物的除草剂
EA201390077A1 (ru) 2010-07-08 2013-06-28 Байер Кропсайенс Нв Белок-переносчик глюкозинолатов и его применения
EP2668278A1 (en) 2011-01-24 2013-12-04 Bayer CropScience NV Use of the rd29 promoter or fragments thereof for stress-inducible expression of transgenes in cotton
EP2524602A1 (de) 2011-05-20 2012-11-21 Bayer CropScience AG Herbizide Mittel für tolerante oder resistente Sojakulturen
UA117816C2 (uk) 2012-11-06 2018-10-10 Байєр Кропсайєнс Акцієнгезелльшафт Гербіцидна комбінація для толерантних соєвих культур
DE202014101590U1 (de) 2014-04-03 2014-04-29 Igus Gmbh Führungssystem für Versorgungsleitungen und Roboter mit Führungssystem

Family Cites Families (1)

* Cited by examiner, † Cited by third party
Publication number Priority date Publication date Assignee Title
DE3765449D1 (de) * 1986-03-11 1990-11-15 Plant Genetic Systems Nv Durch gentechnologie erhaltene und gegen glutaminsynthetase-inhibitoren resistente pflanzenzellen.

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DK437887D0 (da) 1987-08-21
JPH09107981A (ja) 1997-04-28
DK437887A (da) 1988-02-24
ES2038631T3 (es) 1993-08-01
FI873610L (fi) 1988-02-24
JP2815847B2 (ja) 1998-10-27
JPS6371183A (ja) 1988-03-31
JP2749424B2 (ja) 1998-05-13
GR3005200T3 (enrdf_load_stackoverflow) 1993-05-24
IL83604A0 (en) 1988-01-31
CN1040772C (zh) 1998-11-18
HU217208B (hu) 1999-12-28
NZ221526A (en) 1989-03-29
DK175254B1 (da) 2004-07-26
AU7731887A (en) 1988-05-19
JPH0797994B2 (ja) 1995-10-25
JPH07147985A (ja) 1995-06-13
CN87105764A (zh) 1988-11-30
JPH03103193A (ja) 1991-04-30
HUT44621A (en) 1988-03-28
FI100251B (fi) 1997-10-31
CN1124347C (zh) 2003-10-15
EP0257542A2 (de) 1988-03-02
IL83604A (en) 2004-02-19
CA1337597C (en) 1995-11-21
CN1225393A (zh) 1999-08-11
ATE75776T1 (de) 1992-05-15
AU604743B2 (en) 1991-01-03
FI873610A0 (fi) 1987-08-20
EP0257542A3 (en) 1990-03-07

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